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Environmental and Experimental Botany 176 (2020) 104085 Environmental and Experimental Botany 176 (2020) 104085 Contents lists available at ScienceDirect Environmental and Experimental Botany journal homepage: www.elsevier.com/locate/envexpbot Potassium fertilization increases hydraulic redistribution and water use efficiency for stemwood production in Eucalyptus grandis plantations T Verónica Asensioa,*, Jean-Christophe Domecb,c, Yann Nouvellond,e, Jean-Paul Laclaud,e, Jean-Pierre Bouilletd,e, Lionel Jordan-Meilleb, José Lavresa, Juan Delgado Rojasf, Joannès Guillemotd,e,g, Cassio H. Abreu-Juniora a Universidade de São Paulo-Centro de Energia Nuclear na Agricultura (USP-CENA), CEP 13400-970 Piracicaba, SP, Brazil b Bordeaux Sciences Agro, INRA UMR 1391 ISPA, F-33170 Gradignan, France c Nicholas School of the Environment, Duke University, Box 90328, Durham, NC 27708, USA d Eco&Sols, Univ Montpellier, CIRAD, INRA, IRD, Montpellier SupAgro, 34060 Montpellier, France e CIRAD, UMR Eco&Sols, 34060 Montpellier, France f Agro Ambiência Serviços Agrícolas, Piracicaba, SP, Brazil g Department of Forest Sciences, Universidade de São Paulo-“Luiz de Queiroz” College of Agriculture (USP-ESALQ), CEP 13418-900 Piracicaba, SP, Brazil ARTICLE INFO ABSTRACT Keywords: Climate change is expected to increase the frequency of droughts in most tropical regions in the coming decades. Eucalyptus A passive phenomenon called hydraulic redistribution (HR) allows some plant species to take up water from Hydraulic lift deep and wet soil layers and redistribute it in the upper dry layers where other plants and soil biota can benefit Soil fertilization from it. In addition, soil fertilization, particularly potassium (K), may also affect drought-adaptive mechanisms Tree transpiration and increase water use efficiency (WUE) on poor and acidic tropical soils. The present study aimed at quantifying Water use the role of HR and K fertilization on both wood productivity and WUE for stemwood production (WUEp)of Eucalyptus grandis plantations in Brazil under ambient and reduced (−37%) throughfall conditions. Tree transpiration was measured using trunk sap flow sensors over 21 months, and HR was estimated from the reverse sap flow (RF) observed in shallow roots over 18 months. Tree biomass, hydraulic conductance, soil water storage from surface to the water table (down to 17 m), and leaf photosynthetic capacity were also assessed. Significant HR was detected over the whole year, even during the rainy seasons. Neither potassium fertili- zation nor throughfall exclusion affected the velocity of water transported by HR, probably because most trees reached water table. Nonetheless, some photosynthetic capacity parameters, including the maximum photo- synthetic rate (Amax), increased in treatments with K addition. This higher Amax combined with an increased sapwood area index, was associated with an increase in water uptake by 30 %–50 % and WUEp by 300 % relative to K-deficient trees. We postulate that the increase in WUEp promoted by potassium fertilization was partly driven by an increase in biomass allocation to wood, at the expense of foraging organs (leaves and roots), because K addition alleviated constraints on light and water use. Our results indicate that fertilizing E. grandis plantations with K is beneficial to both wood biomass production and WUEp. 1. Introduction and water vapor pressure deficit, and an increase in the length of dry periods (Chou et al., 2012; Salazar et al., 2016). From 2012–2015, the Extreme climatic events such as heavy rainfall, wind storms, and state of São Paulo, in southeast Brazil, suffered a 20–23 % reduction in severe droughts are increasing in frequency worldwide as a con- precipitation, which led to a significant decrease in soil moisture sequence of climate change (IPCC, 2014; Sherwood and Fu, 2014). (Getirana, 2016). The increase in the frequency of extreme climatic Recent climate predictions show that Southeast Brazil will be affected events will increase the difficulty for plants to acclimate physiologically by more frequent heavy rainfall events, higher mean air temperatures and morphologically to the rapidly changing climatic conditions, which ⁎ Corresponding author. Present address: Department of Soil Science, Universidade de São Paulo-“Luiz de QueirozCollege of Agriculture (USP/ESALQ), CEP 13418- 900, Piracicaba, SP, Brazil. E-mail address: [email protected] (V. Asensio). https://doi.org/10.1016/j.envexpbot.2020.104085 Received 30 January 2020; Received in revised form 15 April 2020; Accepted 15 April 2020 Available online 16 May 2020 0098-8472/ © 2020 Elsevier B.V. All rights reserved. V. Asensio, et al. Environmental and Experimental Botany 176 (2020) 104085 will likely result in decreased primary productivity and increased (Stape et al., 2004, 2010; Binkley et al., 2017). Their fast growth is mortality (Choat et al., 2018). Since planted forests play an important often associated with high water-use, thus raising concerns on their role in supplying human needs for wood and paper products, it is im- potential impact on water resources. There is therefore much interest in portant to determine which forest management practices would be assessing their water-use efficiency (WUE), a highly integrative in- more appropriate for trees to grow properly under those new climatic dicator related to the plantation ability to capture carbon and produce conditions (Noormets and Nouvellon, 2015). biomass per unit of water use (Farquhar et al., 1989; Law et al., 2002; High mortality rates in commercial eucalypt plantations during re- Mateus et al., 2019). At the stand level, WUE for stemwood production cent droughts in Brazil (Scolforo et al., 2019a) have highlighted the (WUEp) can be computed as the ratio of stemwood biomass increment need to understand the key physiological processes that allow trees to to the water transpired over the same period. In fact, in areas with acclimate to recurrent dry summers (Lobell et al., 2011; Gessler et al., limited water resources, managing planted forests in a way that in- 2017). A suitable nutrient application, and in particular potassium (K) creases tree WUEp is a promising avenue to sustain productivity under fertilization has been shown to have positive effects on plant water and climate change. light use efficiency during drought periods (Harvey and van den The main goal of our study was to assess the effects of K fertilization Driessche, 1997; Cakmak, 2005; Battie-Laclau et al., 2016). Previous on wood production, water use, hydraulic redistribution between soil studies have evaluated the effect of the interaction between K fertili- layers, whole tree hydraulic conductance (Ktree) and WUEp in Eucalyptus zation and water availability on plant growth, but most of them with grandis plantations grown under ambient and reduced rainfall condi- non-woody species such as corn and sugarcane (da Silva Moura et al., tions, over the three last years of a commercial rotation (from 4 to 6 2005; Djaman et al., 2013; Mahmood et al., 1999, Martineau et al., years after planting). We hypothesized that (i) HR occurs in the studied 2017). Other studies on woody species like olive trees, were performed Eucalyptus grandis plantations, and is more pronounced under reduced on small and potted individuals growing under greenhouse conditions soil water availability; (ii) K addition increases HR and; (iii) K addition (Arquero et al., 2006). Under field conditions, the only studies ad- increases water-use efficiency for stemwood production. While Battie- dressing the potassium nutrition x drought interacting effects on woody Laclau et al. (2016) studied the effects of K fertilization and throughfall species were conducted on Eucalyptus grandis in Brazil (Battie-Laclau exclusion on WUEp over the early growth of the trees in the same ex- et al., 2014; Christina et al., 2015) but were limited to the first three periment, we show in this paper stand water use and WUEp at the end of growing seasons after planting. The end of the rotation cycle (that ty- the rotation cycle, as well as HR and whole-tree water transport (tree pically lasts 6–7 years) is however a crucial period, when most of the hydraulic conductance; Ktree), which were not evaluated in previous drought-related mortality in Brazilian eucalypt plantations occurs studies. (Scolforo et al., 2019b). This is likely due to decreasing soil water availability and increasing water table depth as stand ages (Christina 2. Materials and methods et al., 2018). Potassium fertilization was shown to increase photo- synthesis and tree biomass during the first years after harvest (Battie- 2.1. Experimental design Laclau et al., 2014), but it also increased tree transpiration, thus leading to faster soil water depletion and increased plant water stress during the The experiment was carried out at the Itatinga Experimental Station dry seasons (Battie-Laclau et al., 2014). An evaluation of the interaction of the University of São Paulo in Brazil (23°02´S; 48°38´W). Over the effect of K fertilization and soil water availability on the water use and previous 15 years, the mean annual rainfall was 1360 mm. The mean water-use efficiency of eucalyptus trees along a whole commercial ro- temperature was 15 °C during the dry season (from June to September), tation is thus strongly needed. and 25 °C during the rainy season (from October to May). The experi- Plants have different strategies for surviving during drought per- mental site was located on a hilltop (slope < 3 %) at an altitude of 850 iods. Some plant species such as Eucalyptus trees can take up water from m.
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